RESEARCH ARTICLE New Plants from the Lower Devonian Pingyipu Group, Jiangyou County, Sichuan Province, China Dianne Edwards1,2*, Bao-Yin Geng1, Cheng-Sen Li1 1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing, China, 2 School of Earth and Ocean Sciences, Cardiff University, Cardiff, United Kingdom * [email protected] Abstract a11111 Descriptions of Lower Devonian plants from Yunnan, South China, have revolutionized concepts of diversity and disparity in tracheophytes soon after they became established on land. Sichuan assemblages have received little attention since their discovery almost 25 years ago and require revision. With this objective, fieldwork involving detailed logging and collection of fossils was undertaken in the Longmenshan Mountain Region, Jiangyou County and yielded the two new taxa described here. They are preserved as coalified com- OPEN ACCESS pressions and impressions that allowed morphological but not anatomical analyses. Yan- Citation: Edwards D, Geng B-Y, Li C-S (2016) New menia (Zosterophyllum) longa comb nov is based on numerous rarely branching shoots Plants from the Lower Devonian Pingyipu Group, with enations resembling lycophyte microphylls, without evidence for vasculature. The Jiangyou County, Sichuan Province, China. PLoS ONE 11(11): e0163549. doi:10.1371/journal. presence of sporangia is equivocal making assignation to the Lycopsida conjectural. The pone.0163549 plant was recently described as a zosterophyll, but lacks strobili. These are present in the Editor: Navnith K.P. Kumaran, Agharkar Research second plant and comprise bivalved sporangia. The strobili terminate aerial stems which Institute, INDIA arise from a basal axial complex displaying diversity in branching including H- and K- Received: April 28, 2016 forms. These features characterise the Zosterophyllopsida, although the plant differs from Zosterophyllum in valve shape. Comparisons indicate greatest similarities to the Lower Accepted: September 3, 2016 Devonian Guangnania cuneata, from Yunnan, but differences, particularly in the nature of Published: November 16, 2016 the sporangium border, require the erection of a new species, G. minor. Superficial exami- Copyright: © 2016 Edwards et al. This is an open nation of specimens already published indicate a high degree of endemism at both species access article distributed under the terms of the and generic level, while this study shows that Yanmenia is confined to Sichuan and Guang- Creative Commons Attribution License, which permits unrestricted use, distribution, and nania is one of the very few genera shared with Yunnan, where assemblages also show a reproduction in any medium, provided the original high proportion of further endemic genera. Such provincialism noted in the Chinese Lower author and source are credited. Devonian is explained by the palaeogeographic isolation of the South China plate, but this Data Availability Statement: All relevant data are cannot account for differences/endemism between the Sichuan and Yunnan floras. Such within the paper. an enigma demands further integrated geological, palaeobotanical and palynological Funding: Chinese Academy of Sciences for a studies. Visiting Professorship for Senior Scientists (No. 2013T2S0022 to Dianne Edwards), www.cas.ac. cn. Competing Interests: The authors have declared that no competing interests exist. PLOS ONE | DOI:10.1371/journal.pone.0163549 November 16, 2016 1 / 21 New Lower Devonian Plants from Sichuan, China Introduction Previous studies on the diversification of terrestrial plants in Early Devonian times in China have largely concentrated on the Lower Devonian assemblages of east and south-east Yunnan Province, [1–4]. These indicate exceptional disparity and a high percentage of endemics, many of which defy attempts to elucidate their affinities. The latter include Hsüa robusta [5], Stachyo- phyton yunnanense [6], Catenalis digitalis, Eophyllophyton bellum [7,8], Adoketophyton subver- ticillatum [9], Celatheca beckii [10], Polythecophyton demissum [11] and Bracteophyton variatum [12]. Scrutiny of species lists in Hao and Xue 2013 [4] shows that, of taxa that could be identified with some confidence, only eight genera out of 28 were cosmopolitan (i.e. were found outside the South China Plate) and that four of these were represented by endemic spe- cies (Table 1, this work). Such data hint at the provincialism, as defined by high percentages of endemic taxa, which culminated in the emergence of the late Palaeozoic Cathaysian flora [13], although Xiong et al. 2013 [14] found that the Pragian interval was the only one in the Chinese Devonian in which numbers of endemic genera exceeded those of cosmopolitan ones and that post Pragian proportions of endemic genera declined gradually. By contrast, the Lower Devonian assemblage of Sichuan province, also on the South China plate/palaeocontinent [15], has been little studied since plants were first reported by Li and Cai in 1978 [16] (Sporogonites xichuanensis, Psilophytites) and followed by the descriptions of Geng ([17,18]; Table 1). Exceptions are recent papers by Wang in 2007 [19], who described Zosterophyllum longa (sic) sp. nov., and Xu and Wang in 2009 [20] who queried Geng’s identi- fication of Leclercqia complexa, because the leaves were more frequently branched than in the north American taxon. They did not provide a new name. In view of the recent resurgence of interest in Chinese plants and the availability of new material, a reinvestigation of the Sichuan plants seems long overdue and begins here with the description of two new taxa, including that originally named Zosterophyllum longa by Wang in 2007 [19]. A full species list is given in Table 1, where the legend indicates uncertainties of identification and affinity plus taxa in need of revision. Accuracy of identification, plus confidence in dating, is an essential prerequisite to any inferences on the phytogeographic or evolutionary significance of the assemblage. Table 1. Species list from Geng (1992a, b) and more recently collected fossils. Fossils in lower (L), middle (M) and upper (U) parts of the section. C/E: Cosmopolitan/Endemic, S: Sichuan, G:genus, sp:species, · inaccurate identification. Taxa part Af®nites C/E References ● Eogaspesiea gracilis L Unknown ? Geng 1992a ● Hicklingia cf edwardii L ibid EG Ibid Oricilla unilateralis sp. nov L ibid CG, EspS Ibid ● Psilophyton sp. L Trimerophytopsida? ? Ibid ● Uskiella sp. L Unknown EG Ibid ● Zosterophyllum myretonianum L Zosterophyllopsida ? Ibid ● Z. sichuanensis sp. nov. L ibid EG Ibid Z. yunnanicum L ibid EspChina Ibid Amplectosporangium jiangyouense L Unknown EG Geng 1992b ● Drepanophycus spinaeformis M Lycopsida C Geng 1992a ● D. spinosus M ibid C Ibid Drepanophycus sp M ibid C Ibid ● Leclerqia complexa U ibid ? ibid, Xu et Wang 2007 Guangnangia new sp. U Zosterophyllopsida EspS This paper Yanmenia longa gen. et comb. Nov. L Lycopsida? EG This paper Sciadocillus cuneiformis L Unknown EG Geng 1992a doi:10.1371/journal.pone.0163549.t001 PLOS ONE | DOI:10.1371/journal.pone.0163549 November 16, 2016 2 / 21 New Lower Devonian Plants from Sichuan, China Stratigraphy and Age Deliberations on the age of the plant assemblages in the Pingyipo Group have been hampered by complexity in the geology of the area leading to misunderstandings on stratigraphy, names of strata and correlation. The plant-containing rocks form part of the Tangwangzhai Syncline in the Longmenshan Mountain region where Devonian sequences are found in the east and west limbs in Jiangyou and Beichuan Counties respectively. The most extensive and relatively well-dated exposures are in a stream section between Guixi and Shawozi villages (Guixi-Shawozi Section, Beichuan County) in the west limb with a possibly complete Devonian sequence, 13 km long in a total thickness of 4700m [21]) of which 2064 m comprises the Pingyipu Group. Although fossiliferous, the fishes, ostracodes, bivalves and brachiopods are endemics but based on conodonts, the strata are considered pre-Emsian. Palynological studies indicate that the the Bailiuping Formation above the Pingyipu group is Pragian, while the group itself ranges from probably the Pragian (Guangshanpo Formation) to probably Lochkovian (remaining three formations). Details of the stratigraphy and fossils are given in Table 2. Strata near the base of the succession in the east limb, also called the Pingyipu Group and also overlain by the Bailiuping Formation, are exposed in the Yanmenba Section, Jiangyou County about 60 km from the west limb (Fig 1). The underlying rocks are middle Silurian. However the lithology of the Pingyipu Group and succeeding strata differ in the two limbs such that the formations named in the west cannot be identified in the east. This makes corre- lation, and hence age determinations of the plant fossils, equivocal. The Pingyipu Group in the Yanmenba section is about 672 m thick, considerably less than the same group (2084 m) in the Guixi-Shawozi section. Two sedimentological logs have been Table 2. West limb Guixi-shawozi section. Formation Conodont Invertebrates Fishes zone Upper Maoba Devonian Shawozi Tuqiaozi Middle Guanwushan Devonian Jinbaoshi Lower Yangmaba Devonian Ertaizi Polygnathus serotinus Xiejiawan Ganxi P.perbonus,P. dehiscens Bailiuping Pingyipu Guangshanpo (Br) Orientospirifer cf. wangi Yunnanolepsis cf. chii, Tsuifungshanolepsis Group 243m cf.diandongensis, Yunlongolepis liui, Chuanbeiolepis jiangyouensis, Parapetalichthys